Muscle Transduces Chemical Energy Into Mechanical Energy

Muscle is the major biochemical transducer (machine) that converts potential (chemical) energy into kinetic (mechanical) energy. Muscle, the largest single tissue in the human body, makes up somewhat less than 25% of body mass at birth, more than 40% in the young adult, and somewhat less than 30% in the aged adult. We shall discuss aspects of the three types of muscle found in vertebrates: skeletal, cardiac, and smooth. Both skeletal and cardiac muscle appear striated upon microscopic observation; smooth muscle is nonstriated. Although skeletal muscle is under voluntary nervous control, the control of both cardiac and smooth muscle is involuntary.

The Sarcoplasm of Muscle Cells Contains ATP, Phosphocreatine, & Glycolytic Enzymes

Striated muscle is composed of multinucleated muscle fiber cells surrounded by an electrically excitable plasma membrane, the sarcolemma. An individual muscle fiber cell, which may extend the entire length of the muscle, contains a bundle of many myofibrils arranged in parallel, embedded in intracellular fluid termed sar-coplasm. Within this fluid is contained glycogen, the high-energy compounds ATP and phosphocreatine, and the enzymes of glycolysis.

The Sarcomere Is the Functional Unit of Muscle

An overall view of voluntary muscle at several levels of organization is presented in Figure 49-1.

When the myofibril is examined by electron microscopy, alternating dark and light bands (anisotropic bands, meaning birefringent in polarized light; and isotropic bands, meaning not altered by polarized light) can be observed. These bands are thus referred to as A and I bands, respectively. The central region of the A band (the H band) appears less dense than the rest of the band. The I band is bisected by a very dense and narrow Z line (Figure 49-2).

The sarcomere is defined as the region between two Z lines (Figures 49-1 and 49-2) and is repeated along the axis of a fibril at distances of 1500-2300 nm depending upon the state of contraction.

Myofibril

Figure 49-1. The structure of voluntary muscle. The sarcomere is the region between the Z lines. (Drawing by Sylvia Colard Keene. Reproduced, with permission, from Bloom W, Fawcett DW: A TextbookofHistology, 10th ed. Saunders, 1975.)

Myofibril

Figure 49-1. The structure of voluntary muscle. The sarcomere is the region between the Z lines. (Drawing by Sylvia Colard Keene. Reproduced, with permission, from Bloom W, Fawcett DW: A TextbookofHistology, 10th ed. Saunders, 1975.)

The striated appearance of voluntary and cardiac muscle in light microscopic studies results from their high degree of organization, in which most muscle fiber cells are aligned so that their sarcomeres are in parallel register (Figure 49-1).

Thick Filaments Contain Myosin; Thin Filaments Contain Actin, Tropomyosin, & Troponin

When myofibrils are examined by electron microscopy, it appears that each one is constructed of two types of longitudinal filaments. One type, the thick filament, confined to the A band, contains chiefly the protein myosin. These filaments are about 16 nm in diameter and arranged in cross-section as a hexagonal array (Figure 49-2, center; right-hand cross-section).

The thin filament (about 7 nm in diameter) lies in the I band and extends into the A band but not into its H zone (Figure 49-2). Thin filaments contain the proteins actin, tropomyosin, and troponin (Figure 49-3). In the A band, the thin filaments are arranged around the thick (myosin) filament as a secondary hexagonal array. Each thin filament lies symmetrically between three thick filaments (Figure 49-2, center; mid cross-

section), and each thick filament is surrounded symmetrically by six thin filaments.

The thick and thin filaments interact via cross-bridges that emerge at intervals of 14 nm along the thick filaments. As depicted in Figure 49-2, the cross-bridges (drawn as arrowheads at each end of the myosin filaments, but not shown extending fully across to the thin filaments) have opposite polarities at the two ends of the thick filaments. The two poles of the thick filaments are separated by a 150-nm segment (the M band, not labeled in the figure) that is free of projections.

The Sliding Filament Cross-Bridge Model Is the Foundation on Which Current Thinking About Muscle Contraction Is Built

This model was proposed independently in the 1950s by Henry Huxley and Andrew Huxley and their colleagues. It was largely based on careful morphologic observations on resting, extended, and contracting muscle. Basically, when muscle contracts, there is no change in the lengths of the thick and thin filaments, but the H zones and the I bands shorten (see legend to Fig-

A. Extended

H band

I band

A band

Z line

2300 nm

Actin filaments 6-nm diameter

Cross section:

2300 nm

Actin filaments 6-nm diameter

Cross section:

a-Actinin

Myosin filaments 16-nm diameter a-Actinin

Myosin filaments 16-nm diameter

B. Contracted c

Thin-filament

Thick — H filament

1500 nm

6-nm diameter

Figure 49-2. Arrangement of filaments in striated muscle. A: Extended. The positions of the I, A, and H bands in the extended state are shown. The thin filaments partly overlap the ends of the thick filaments, and the thin filaments are shown anchored in the Z lines (often called Z disks). In the lower part of Figure 49-2A, "arrowheads," pointing in opposite directions, are shown emanating from the myosin (thick) filaments. Four actin (thin) filaments are shown attached to two Z lines via a-actinin. The central region of the three myosin filaments, free of arrowheads, is called the M band (not labeled). Cross-sections through the M bands, through an area where myosin and actin filaments overlap and through an area in which solely actin filaments are present, are shown. B: Contracted. The actin filaments are seen to have slipped along the sides of the myosin fibers toward each other. The lengths of the thick filaments (indicated by the A bands) and the thin filaments (distance between Z lines and the adjacent edges of the H bands) have not changed. However, the lengths of the sarcomeres have been reduced (from 2300 nm to 1500 nm), and the lengths of the H and I bands are also reduced because of the overlap between the thick and thin filaments. These morphologic observations provided part of the basis for the sliding filament model of muscle contraction.

Diabetes 2

Diabetes 2

Diabetes is a disease that affects the way your body uses food. Normally, your body converts sugars, starches and other foods into a form of sugar called glucose. Your body uses glucose for fuel. The cells receive the glucose through the bloodstream. They then use insulin a hormone made by the pancreas to absorb the glucose, convert it into energy, and either use it or store it for later use. Learn more...

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